Sandalwood Research Newsletter

February 2003 Issue 17 ISSN 1321-0022X

EDITOR’S NOTE

This edition of the SRN examines host trials with quandong (Santalum acuminatum) and genetic variation within Indian sandalwood (Santalum album). In South Australia, S. acuminatum is being grown in plantations for its edible red fruit. Matthew Lethbridge is examining S. acuminatum performance with different Acacia species that also have the potential for commercial use in the wattle seed industry. In previous trials, Acacia species have proven to be very good hosts for S. spicatum and S. album.

Studies on S. album from India and West Timor indicate that there is a high degree of variation within the species, and there may in fact be separate varieties or races. Within India, Angadi, Jain and Shankaranarayana examine the levels of genetic diversity within and between eight populations of S. album. Jain, Angandi and Shankaranarayana also discuss environmental, morphometric and genetic characteristics of nine S. album populations growing within India.

Recently I changed Departments, and I now work for the Forest Products Commission, Western Australia. However I am still researching sandalwood and I will continue to be editor of the SRN. My new mailing address is shown at the end of this newsletter.

Jon Brand

acacias as host plants for quandong Progress report: Integrated wattle and quandong orchard fruit production.

Matthew Lethbridge Materials and Methods Australian Quandong Industry Association (AQIA), P.O. Box 1160, Loxton, South Australia, 5333 Within the research area of the Aus- tralian Arid Land Botanic Gardens is Abstract a chenopod heathland (predominantly Eight species of semi arid acacia species with potential for wattle seed produc- Atriplex and Maireana species, these tion in the native food industry were tested for there suitability as hosts for quan- were retained) of deep alkaline (pH dong (Santalum acuminatum) in an irrigated area at the Australian Arid Lands 9.5) loam of zero inclination. The Botanic Gardens, Port Augusta, South Australia. Preliminary results indicate area had been previously deep ripped that Acacia victoriae is superior to the seven other species tested. (1992). Eight species of semi-arid acacia (Table 1) were planted in a grid pattern of 6 m by 4 m. Each species Introduction A survey by Maslin et al (1998) iden- was planted as two rows of 11 plants tified 47 species of acacia that have from tube stock grown on site. They The quandong fruit (Santalum acumi- potential for both seed production for were irrigated with single drippers (8 natum) and wattle seed from acacia human consumption and cultivation in litres per hour) at the base of each species occupy significant niches in the southern semi arid regions of acacia for 2hrs, twice a week (32 li- the fledgling native food industry Southern Australia. There has been a tres per week). Acacia papyrocarpa, (Graham and Hart 1996, Ahmed and small but increasing demand for wat- A. victoriae and Santalum acumi- Johnson 2000). Acacia species are tle seed in the native food industry natum occur naturally on the site showing significant promise in the and is part of investigations (all be it (outside the irrigated area). sandalwood industry as host plants, minor) into the commercial output in (Brand et al., 2001, Radomiljac et al., the broad scale planting of acacias to Procedure 1999a) with the expectation that legu- combat dryland salinity (Simpson and In September 1999, the acacia seeds minous species could also provide Chudleigh 2001). The current study were germinated and transplanted to valuable timber products. is extending the theme of utilising the large forestry tubes. At age seven multipurpose nature of leguminous

inside... Progress report: Integrated wattle and quandong orchard Pages 1-4 Genetic diversity between sandal populations …. in India Pages 4-5 Edaphic, environmental and genetic factors associated with growth … Pages 6-7 months, the acacia seedlings were Table 1. The Eight acacia species and their seed source, selected as hosts for planted with a handful of slow release S. acuminatum. The species were selected from the list prepared by Maslin native Osmocote and mulching. (1998) and suitability for the site. There was no further addition of fer- tiliser to the site. In April 2001, two Species Source germinated kernels of quandong (orchard derived seed, Whyalla South A. murrayana 70 km. West of Cobar (AALBG collection) Australia) were direct seeded within A. victoriae Kenmore Park SW (thornless) (AALBG) ten centimeters of each dripper. Quandong heights were recorded in A. rivalis RG 554 K (Australian Bush products) April 2002 and October 2002 (Figures A brumalis CALM D1001 1 and 2). A calamifolia 44 Km. West of Yunta (AALBG) Statistics A hakeoides Quorn (AALBG) Quandong mean heights were com- pared using one-way ANOVA, and A hemiteles CALM N98177 Least Significant Difference (LSD) A. argyrophylla Carrieton (AALBG) tests.

of quandong seedlings can be prob- victoriae is currently the most impor- Results lematical. The most economic and tant wattle used in the Australian Na- efficient method for establishing S. tive Food Industry (Maslin 1998), and It is evident from the results (Table 2) spicatum is by direct seeding. (Brand is likely to continue so, given its po- that Acacia victoriae appears to be a and Jones 1999). This methodology tential dual role. Native food growers superior host to the seven other se- may have practical use in the quan- have used this combination because of lected native food acacias. None of dong industry. its native food association and proba- the quandongs showed any obvious bly more as a legume of convenience nutrient deficiency symptoms by vis- than from scientific rational ual leaf inspection. (see Barrett et al “Acacia victoriae is currently the (Schwarz, 2001). 1993). It was also noted that where most important wattle used in the two germinants survived, both plants Australian Native Food Industry “ A significant feature of Acacia victo- usually showed similar heights, per- riae is its high salt tolerance (Maslin haps indicating the vital role of the Discussion 1998). Santalum acuminatum is clas- host acacia’s root system in sustaining sified as a salt tolerant plant (Walker growth of the quandong. It should Quandongs planted near Acacia victo- 1989). Hence this combination may also be noted that the quandong sur- riae showed significantly better have value in salt land reclamation. vival from germinated seed was growth characteristics for the first 18 Neither species has yet been recog- 53 ± 4.5 % across the eight species, months of growth when compared to nised for this purpose (Marcar 1995). and for Acacia victoriae 50 % of the the other seven host species at this It is also reasonable to assume that planted germinated seed survived. It site. From a commercial point of studies with other combinations of is often reported that transplantation view this is a good result as Acacia acacia/ santalum may have some pre-

(a). (b).

Figure 1ab. Quandong, age 18 months, growing with 30 month old (a) Acacia victoria—quandong ht: 1.24 m; and (b) Aca- cia argophylla—quandong ht: 1.1 m. Table 2. Growth parameters of quandong (18 month old) associated with indi- Barrett D. and Fox J., (1993). Santa- cated acacia species (30 months old). Same letter following score indicates that lum album. Photographic record of they are not significantly different (LSD 0.05). nutrient deficiency symptoms in seedlings Aisuli 6, 1993 Species Average Quandong Brand, J.E., Crombie, D.S. and ht. (cm.), October 2002 Mitchell, M.D. (2000). Establish- ment and growth of sandalwood A. victoriae 98.2 a (Santalum spicatum) in south- A. hemiteles 81.0 b western Australia: the influence of host species. Australian Forestry A. argyrophylla 74.7 bc 63 (1): 60-65. A. murrayana 68.6 bc Brand, J.E., Fox, J.E.D. and Moretta, P. (2001). Review of research into A. calamifolia 66.2 cd sandalwood (Santalum spicatum) A. rivalis 62.0 cd tree farm systems in south-western Australia. In Conference Proceed- A. brumalis 58.2 cd ings: Forests in a Changing Land- A. hakeoides 52.6 d scape: 16th Commonwealth For- estry Conference jointly with the 19th Biennial Conference of the dictive value in species selection. A ductions in growth. A study of the Institute of Foresters of Australia, S. album host study by Radomiljac water use rates of the eight species Fremantle, Western Australia, 18- (1999a) included Acacia ampliceps, used in this study and others may 25 April, 2001 Promaco Conven- which has high salt tolerance (Marcar yield some useful correlations with tions, Perth, pp. 527-535. 1995) and out performed Acacia the hemi parasites growth rate. A Brand J. and Jones P. (1999). Grow- trachycarpa as a host for tropical san- host study by Brand et al (2000) con- ing Sandalwood (Santalum spica- dalwood. Another pot study by Byrne cluded that the faster growth rate of S. tum ) on farmland in Western Aus- (1998) used the highly salt tolerant spicatum near A. acuminata did not tralia. Sandalwood Information A. cyclops (Marcar 1995) but could appear to be due to greater access to sheet No 1. CALM. not recommend this species as a good water, especially during the summer Bryne B. (1998). The Physiology and host. period but this study compared Aca- Biochemistry of the interaction be- cia, Allocasuarina and Eucalyptus tween the root hemiparasite Santa- Acacia brumalis has sometimes been species, where nutritional differences lum acuminatum and its host plants. found growing in moderately saline may be more significant. Ph. D. Thesis, Flinders University soils (Maslin 1998), but in this study Host : parasite interactions are clearly (South Australia). it did not stand out as a host for quan- complex and empirical data will al- Graham C., Hart D. (1997). Bush- dong. Similarly A. saligna is often ways be a valuable means of assess- food. In: The New Rural Industries. found on moderately saline soils ing species as host plants. A handbook for Farmers and Inves- (Marcar 1995, Maslin 1998). In a tors. Edited by Hyde K. RIRDC similar study (although non irrigated) Marcar N. Crawford D., Leppert P., in WA, Brand et al (2001) found Acknowledgements. Jovanovic T., Floyd R., Farrow R., A. saligna to be a very good primary (1995) Trees for Saltland. A guide host for Santalum spicatum, when The financial support of the Rural to selecting native species for Aus- compared to A. acuminata, Industries Research and Development tralia. CSIRO Australia A. hemiteles and A. microbotrya. Corporation (RIRDC) and the Austra- Maslin B., Thomson L., McDonald However, this may be attributable to lian Quandong Industry Association. M., Hamilton-Brown S. (1998). the fast growth rate of this species, (AQIA) is gratefully acknowledged. Edible wattle seeds of Southern nevertheless, combinations of host The technical help of Bernie Haas and Australia. A review of species for and quandong with revegetation value other staff at the Australian Arid Land use in semi-arid regions. CALM in saline areas can only be considered Botanic Gardens (AALBG) has been and CSIRO Australia. a useful addition to farm management most helpful. Thanks to Perce Radomiljac A., McComb J., Pate J. practices. The attribute of salt toler- Prenzel for the gift of quandong seed (1999a). Heterotrophic carbon gain ance in plants is often associated with and Graham Herde for his hospitality and mineral nutrition of the root water conservation strategies. and useful discussion. hemi-parasite Santalum album in pot culture with different hosts. Little or no research has been done at Australian Forestry 62 92): 128- this stage into the rate of water use of References 138 acacia species (Simpson and Chud- Radomiljac A., McComb J., Pate J. leigh 2001). Species with high water Ahmed A. and Johnson K. (2000). (1999b). Gas exchange and water use rates, will perpetuate a less fa- Turner Review No 3. Horticultural relations of the hemi-parasite San- vourable water potential gradient development of Australian native talum album in Association with (Radomiljac 1999b, Byrne 1998) from edible plants. Aust. J. Bot. 48: 417- legume and non-legume hosts. An- host to parasite with consequent re- 426. nals of Botany 83:215-224 Walker R., (1989). Growth, photo- used for isoenzyme analysis and re- Scharwz K. (2001). Australian Quan- synthesis and distribution of chlo- sults obtained (Angadi et al) were dong Industry Conference Field trip ride, sodium and potassium ions in used to study the genetic variation. A Loxton 2001. salt affected quandong (Santalum method of Nei (1972 and 1973) which Simpson S. and Chudleigh P. (2001). acuminatum). Aust. J. of Plant is based on identity of genes between Wattle seed production in Low Physiology 16: 365-377 populations was used for the purpose. Rainfall Areas. RIRDC publication Of the eight enzymes tested, four en- no. 01/08 zymes namely POD, MDH, GDH, GLUDH were active in the seed tis- sue. Genetic diversity between sandal populations of different provenances in India Results and Discussion Angadi. V.G., Jain, S.H. and Shankaranarayana, K.H. The mean genetic diversity measures Institute of Wood Science and Technology, Malleswaram, Bangalore, 560003, for Santalum album L. populations of India. different provenances are presented in Table 1. The mean number of alleles Abstract per locus ranges from 1.4 (Bangalore Genetic distance between Sandal (Santalum album L.) populations of different and Seoni) to 3.6 (Mandagadde). The provenances has been studied, using isoenzyme data. Results show that there is a percentage of polymorphic loci per gap in genetic distance between eight identified provenances of Sandal which population varies from 33.0% (Seoni) indicates that they are of separate varieties or races, thus further confirming their to 69.2% (Marayoor). The maximum status. heterozygosity (0.50) was found in Bangalore provenance and minimum (0.18) in Mandagadde provenance. Introduction normalized identity of genes between Other provenances show medium populations is equivalent to protein heterozygosity (0.26 to 0.44). Sandal, which is represented in the identity. The term genetic distance is genus Santalum, is indigenous to pen- then defined as number of nucleotide Table 2 indicates estimates of Nei’s insular India. Earlier it was of the or codon differences per unit length of (1972) genetic distance (D) between opinion that Sandal was introduced to DNA molecule. Sandal populations of different prove- India from Timor, in Indonesia nances. Normalized genetic identity (Fischer, 1928, 1938). A recent study between Koraput and Mandagadde by Brand (1994) concluded that San- “Thus the gap in genetic distance be- and between Koraput and Javadi were dal in West Timor and India are of tween different provenances of Sandal both close to zero, hencfe calculation different varieties or races, because of indicates that they are separate varie- of genetic distance between them large genetic distance between the ties or races “ were omitted (Nei, 1972). Genetic two countries. distance between Mandagadde and Materials and Methods the Bangalore provenance is found to In India, Sandal is distributed all over be minimum (0.125), where as genetic the country and more than 90% of it Santalum album L. seeds were col- distance between Thangli and Seoni- lies in Karnataka and Tamil Nadu. lected from eight different provenance provenance is found to be maximum Based on a recent survey undertaken areas during fruiting season. Twenty (0.585). in India, eight potential Sandal bear- individual seeds per population were ing areas in different eco-climatic zones have been identified as poten- Table 1. Mean genetic diversity estimates (Nei’s 1972) of different provenances tial provenances (Jain et al 1997) and of sandal. their status was confirmed using isoenzyme study (Angadi et al, in press). In spite of their known ge- S. No Population Sample Mean No of % of Poly- Heterozygosity netic identity with reference to vari- Size Alleles per morphic (figures in brackets ous parameters, information on their locus Loci indicate stand. error) genetic diversity between sandal 1 Mandagadde 20 3.6 57.1 0.18 (0.14) populations of different provenances 2 Bangalore 20 1.4 37.5 0.50 (0.23) is lacking. In this present work, at- 3 Thangli 20 1.6 33.5 0.44 (0.15) tempt has been made to study the ge- netic distance between Sandal popula- 4 Marayoor 20 2.7 69.2 0.26 (0.25) tions of different provenances using 5 Chitteri 20 1.8 55.6 0.42 (0.20) isoenzyme study. The concept of 6 Javadi 20 1.5 40.0 0.38 (0.28) genetic distance used here is based on Nei’s (1971) theory which means 7 Seoni 20 1.4 33.0 0.43 (0.21) 8 Koraput 20 1.8 50.0 0.40 (0.22) Table 2. Genetic Distance (D) between sandal populations of different prove- Fischer, C.E.C (1938). Where did the nances (using all loci). Sandalwood tree (Santalum album L) evolve? J. Bombay Nat. Hist. Population Man Ban Mar Seo Tha Chi Jav Kor Soci. 40: 458-460. Jain, S.H., Angadi, V.G., Rajeevalo- Man ------chan, A.N., Shankarnarayana, Ban 0.125 ------K.H., Theagarajan, K.S. and Ran- gaswamy, C.R. (1998). Identifica- Mar 0.131 0.161 ------tion of Provenances of Sandal in Seo 0.161 0.260 0.174 ------India for genetic conservation. In: Radomiljac, A.M., Ananthapadma- Tha 0.252 0.260 0.357 0.585 ------nabho, H.S., Welbourn, R.M. and Chi 0.292 0.168 0.337 0.276 0.328 ------Satyanarayana Rao, K. (Eds.), Proceedings of an international Jav 0.292 0.357 0.456 0.420 0.482 0.432 ------seminar held on 18-19 December Kor - 0.482 0.561 0.469 0.509 0.328 ------1997, Bangalore, India. ACIAR Proceedings No. 84:117-120.

Table 3. Mean Genetic Distance (D) between individual provenance and other Nei, M., (1971). Interspecific gene identified provenances. differences and evolutionary time estimated from electrophoretic SI. No. Provenance Mean Genetic Distance (D) from other data on protein identity. Ameri- Provenances can Naturalist 105:385-398. 1 Mandagadde 0.55 Nei, M, (1972). Genetic distance be- 2 Bangalore 0.26 tween populations. American Naturalist 106: 283-292. 3 Thangli 0.31 Nei, M., (1973). Analysis of gene diversity in subdivided popula- 4 Marayoor 0.34 tions. Proceedings of the National 5 Chitteri 0.40 Academy of sciences of the USA 70: 3321-3323. 6 Javadi 0.31 7 Seoni 0.49 8 Koraput 0.85

Data in Table 2 were used to calculate that they are of separate varieties or mean genetic distance between indi- races. The larger genetic distance vidual provenances and other identi- between them further confirms their fied seven provenances, and are repre- status. And it is also concluded that sented in Table 3. From this, it is the provenance of Bangalore was found that Bangalore and other prove- found to be the best in terms of het- nances had a minimum mean genetic erozygosity and hence suitable for distance of 0.26; Koraput and other breeding and propagation work on provenances had a maximum mean Sandal. genetic distance of 0.85. The mean genetic distance between other prove- nances ranges from 0.31 (Marayoor / Acknowledgements Chitteri and other provenances) to 0.55 (Mandagadde and other prove- The authors are grateful to Dr K.S nances). Thus the gap in genetic dis- Rao, Director of the Institute, for his tance between different provenances interest and encouragement during the of Sandal indicates that they are sepa- course of this study. rate varieties or races which further confirms their status. References

Brand, J.E. (1994). Genotypic varia- Conclusion tion in Santalum album. Sandal- wood Reasearch Newsletter 2: 2-4. The genetic diversity measures for Fischer, C.E.C (1928). The original Santalum album L. population of dif- home of Santalum album L., J. ferent provenances very well indicate Indian Bot. Soc. 7: 12-13. Edaphic, environmental and genetic factors associated to vary from 0.052 to 0.292. Thus the larger genetic distance between differ- with growth and adaptability of Sandal (Santalum album ent provenances concludes that they L.) in provenances are of separate varieties or races.

The observed variation in trees from Jain, S.H., Angandi, V.G. and Shankaranarayana, K.H. widely different parts of the species range could be referred to as geo- Institute of Wood Science and Technology, Malleswaram, Bangalore, 560003, graphical or racial variation. From India. the history, it is known that the sandal seeds are originally either spread Abstract through natural selection or by means Sandal tree grows under different edaphic and eco climatic conditions. Consid- of propagation from central prove- ering large genetic distance between provenances, it is concluded that under di- nance of Mysore State. Over a period verse locality factors sandal adapts very well in terms of growth, heartwood and of years these trees might have ad- oil content. justed to local environmental and ed- aphic factors leading to the establishe- ment of geographical races or prove- nances in this species, which has a and longitude and eco-climatic (Jain Introduction large genetic distance. The existence et al 1998). Status of these prove- of different provenances within spe- nances were confirmed by isoenzyme Nearly 90% (8100 sq.km) of the total cies may be taken into consideration studies (Angadi et al 1999), which area of Sandal (Santalum album L.) in while making a choice of planting also found that the girth of sandal tree India is in Karnataka and Tamil Nadu. material for further multiplication and influences the percentage of oil. The rest is distributed in most other mass propagation. It is seen from Heartwood from young trees (around States: Andhra Pradesh, Kerala, Table 1 that, sandal adapts very well 10 years of age, height <10m, girth Madhya Pradesh, Orissa, Maharash- under diverse locality factors in terms <50 cms) contains 0.2 to 2% of oil tra, Rajasthan, Uttar Pradesh, Bihar of growth, heartwood, oil content and and that from the mature trees (30 to and Manipur. biochemical characteristics. 50 years of age, height 20 m, girth

100 cms) contains 2.8 to 6.2% of oil. Sandal tree is mainly exploited for its heartwood which yields the renowned Acknowledgements East Indian Sandalwood oil, rated “The existence of different prove- very high for its sweet fragrant, per- nances within species may be taken The authors are thankful to Dr. K.S. sistent, spicy, warm, woody note, into consideration while making a Rao, Director of the Institute for his tenacious aroma and fixative property. choice of planting material for further encouragement during the course of Sandal heartwood currently priced at multiplication and mass propagation” this study. Rs.12 lakhs per ton and its oil Rs,

22,000/-per kg.

At present, India produces 1000 tons References of heartwood and 40 tons of oil per In Table 1, data in respect of area, annum. For many years till now, san- density, latitude, longitude, altitude, Angadi. V.G., Jain, S.H. and dalwood oil alone contributed nearly rainfall, temperature, soil type, pH, Shankaranarayana, K.H. (2003). 25% of the revenue earned from the percentage of oil and genetic distance Genetic diversity between sandal export of various essential oils. (Angadi et al. 2003), from all nine populations of different prove- sandal provenances is shown. nances in India. Sandalwood Re- The aim of this provenance investiga- search Newsletter 17: 4-5. tion is to gather improved knowledge Angadi, V.G., S. Ramalakshmi, S.H. of genetic variation of the species. It Results and Discussion Jain, C.R. Rangaswamy and K.S. is therefore, necessary that the pur- Theagarajan (1999). Allozymic pose of provenance testing should The provenances Bangalore, variation in the seed tissue of San- combine both genetic variability and Marayoor and Thangli (Table 1) are dal (Santalum album L.) population environmental factors. growing well at an altitude of 1000m, of different provenances. average annual rainfall 850-1450 mm, Jain, S.H., V.G. Angadi, A.N. Ra- o max. temperature 33-38 C, minimum jeevalochan, K.H. Shankaranara- o Material and Methods temperature 8-12 C, on soil red sandy yana, K.S. Theagarajan and C.R. Rangaswamy (1998). Identification loam or black clay having slightly of provenances of Sandal in India Nine sandal bearing areas within India acidic to slightly basic pH (6.5 to 7.5) for genetic conservation. ACIAR were identified as potential prove- and moderate nutrient status. The Proceedings, No.84, 117-120, 1998. nances on the basis of population den- genetic distance (D) existing between Jain, S.H. and C.R. Rangaswamy sity, phenotypic character, latitude, populations of provenances was found (1998). Soil properties and their relationship to the growth of Sandal tion Sandal (Santalum album L). A.N. Rajeevalochan , K.S. Theaga- (Santalum album L) in three study FAFAI Journal, 49-53, July-Sept. rajan, C.R. Sarma and C.R. Rangas- areas in Karnataka. Myforest 1999. wamy (1997). A rapid method of 24:141-146. Rangaswamy, C.R., S.H. Jain and K. estimating essential oil content in Jain S.H., V.G. Angadi, G. Raviku- Parthasarathy (1986). Soil proper- heartwood of Santalum album Linn. mar, K.S. Theagarajan and K.H. ties of some Sandal bearing areas. Current Science 72(4): 241-242. Shankaranarayana (1999). Studies Van Vigyan 24 (3&4): 61-68 on cultivation and chemical utiliza- Shankaranarayana K.H., V.G. Angadi,

Table 1. Edaphic, environmental and genetic factors of Sandal provenances.

Prove- Forest Sandal Density Latitude Altitude Mean Temp. Soil pH % of Girth Genetic Genetic nance Division bearing trees/ha Longitude Annual Max. Type oil range identity distance State area ha. in area Rainfall Min. (cm) mm ˚C Bangalore Banga- 24648.0 500 12˚58' N 1000 m 850 36.8 Red loam 6.3-6.5 3.19 30-140 0.75 0.125 lore Kar- 0 77˚ 38' E 12.2 Acidic nataka Thangli Chicka- 47121.0 300 13˚ 40' N 766 m 1500 44.0 Red loam 7.5-7.8 2.76 30-100 0.56 .052 magalur 0 76˚ 00' E 10.5 & Allu- Alkaline Karna- vium taka Manda- Shimoga 65529.0 100 13˚ 9' N 650 m 2000 38.1 Red loam 5.5-5.8 1.55 30-70 * * gadde Karna- 0 75˚ 40' E 13.0 Acidic taka Chitteri Harur 60000.0 70 12˚ 0' N 1050 m 1000 35.2 Red 6.0-6.3 2.06 30-60 0.51 0.292 Tamil 0 78˚ 6' E 8.2 sandy Acidic Nadu loam Javadis Tirupat- 16517.0 90 12˚ 3' N 930 m 1200 38.0 Red loam 6.6-6.7 1.59 30-70 0.51 0.292 tur Tamil 0 78˚ 7' E 12.4 Acidic Nadu Marayoor Munnar 1496.67 500 10˚ 1' N 1000 m 1450 36.0 Black 6.2-6.7 3.15 30-100 0.74 0.131 Kerala 77˚ 1' E 10.0 clay Acidic Koraput Rayagad 2540.00 200 19˚ 55' N 859 m 1525 38.0 Red 6.2-6.6 0.8 30-50 0.0025 - Orissa 82˚ 35' E 4.5 sandy Acidic loam Seoni Seoni 3000.00 60 22˚ 1' N 900 m 1600 40.0 Laterite 6.4-6.9 2.06 30-80 0.69 0.161 Madhya 79˚ 5' E 5.0 Acidic Pradesh Horsely Chittoor 3050.00 70 13˚ 5' N 4312 m 900 30.0 Red loam 6.5-6.8 3.34 30-80 # # Hills East 78˚ 8' E 5.0 Acidic

* Genetic identity / Genetic distance measured from Mandagadde provenance # Newly identified provenance work is in progress

The Sandalwood Research Newsletter (SRN) is published by the Forest Products Commission Authors contact details: and is distributed free of charge. It is intended as a forum for information exchange Matthew Lethbridge on Santalum species worldwide. Articles on a range of Santalum species research Australian Quandong Industry and management issues are welcomed by the Association (AQIA) Sandalwood Research Newsletter. P.O. Box 1160, Loxton, South Australia, 5333 If you would like to contribute an article to the SRN Australia or wish to be included on the SRN mailing list, Tel: + 61 8 8383 6181 please send details to the Editor stating: Fax: + 61 8 8383 6181 your name, title, position, organisation, Email: [email protected] postal address, telephone, fax and email address.

K.H. Shankaranarayana Editor: Jon Brand Forest Products Commission Institute of Wood and Technology MidWest Sharefarms MALLESWARAM Lot 1 / 260 Kalamunda Rd BANGALORE 560003 South Guildford WA 6055 INDIA Australia Tel: + 091 80 334 1731 Fax: + 091 80 334 0529 Telephone: + 61 8 9279 4088 Email: [email protected] Facsimile: + 61 8 9279 5481 Email: [email protected]

The Sandalwood Research Newsletter is produced by the Forest Products Commission, Western Australia.